In studies of heterochrony, timing of development is generally measured relative to some intrinsic dimension, usually size, which serves as a proxy for time. A complementary approach is to measure timing relative to chronological time, an extrinsic dimension. Here, colony growth of a marine hydroid, Hydractinia echinata, is used to compare size- and age-based approaches to rate heterochronies. Colonies consist of feeding polyps; a basal, encrusting mat; and, in some cases, stolons which extend from the mat. Size is measured by the area covered by the mat and, if present, the stolons. Rate of polyp production was compared between colonies using clonal replicates grown in a common environment. Age-based comparisons employed polyp specific growth rates calculated over the same ontogenetic time interval for each replicate. Specific growth rates for colony area were also calculated; rates-ratios (polyp/area) provided size-based measures of polyp growth, analogous to allometric coefficients commonly used in studies of rate heterochronies. These comparisons show that size can be a misleading referent for between-colony comparisons because the rate of areal growth is not constant among colonies. Size-based measures of changes in developmental rates may provide insight into patterns, but not necessarily processes, of heterochrony. The implications of the pattern-oriented framework of size-based heterochrony should be considered in paleontological studies.

Several very large, taxonomically standardized data sets have been compiled and utilized to investigate biogeographic and evolutionary patterns of continental margin benthic foraminifera. Mean partial species durations for 87 frequently occurring and 180 rarely occurring species on the Atlantic continental margin of North America are the same, namely 21 m.y. The global fossil record of these species indicates no center or centers of origin and indicates very rapid dispersal. The Miocene had the greatest number of first occurrences with 46%, followed by the Pleistocene, Pliocene and Oligocene with approximately 13% each. The remaining 14% first occur in the Eocene, Paleocene, and Cretaceous. Species with a wide geographic distribution often exhibit longer species durations than those with narrow geographic ranges. The vast majority of endemic species (150 of 175) occur rarely and have no fossil record.

The Mesozoic marine revolution theory postulates a causal connection between the Cretaceous radiations of durophagous predators and the decline of suspension-feeding echinoderms in shallow-water habitats. In order to test the temporal distribution of dense ophiuroid populations for such a decline, I present a method of calculating the expected distribution of populations or communities in different geologic timespans. This statistical null hypothesis may then be compared with data from the fossil record to draw paleoecological inferences. The model takes into account the relative lengths of time blocks, the decay of sedimentary rock, and changes in shallow sea area through time. Although mass extinctions did not cause the immediate disappearance of shallow-water “brittlestar beds,” brittlestar beds show a significant decline in the Early Cretaceous. Results of several studies suggest that predators were at least partially responsible for reducing the distribution of dense ophiuroid populations.

An investigation of shell morphometrics of the prosobranch genus Paramelania from Lake Tanganyika shows striking contrasts between some live and dead populations from identical localities. Living populations and surficial dead shells were collected from 15 sublittoral-profundal localities along the east side of the lake. Interpopulation variability in this endemic gastropod is clinal (N-S) for several shell characters. Intrapopulation variability of dead shell populations frequently exceeds that of live populations. Distinctive morphs may be present in a local dead shell population which are absent in the live population from the same locality. However, the phenodeviant dead shell morphology may occur among live snails elsewhere in the lake. Phenodeviant shells may be encrusted, but are unabraded and show no preferred orientation.

Radiocarbon ages of >2000 yr b.p. on dead phenodeviants suggest that surficial shell accumulations in Lake Tanganyika are strongly time-averaged. Although wave activity and biogenic concentration are important processes in Lake Tanganyika, neither is sufficient under modern lake conditions to account for these admixed assemblages. Winnowing and in situ stratigraphic condensation during periods of lowered lake levels is the most likely explanation for the occurrence of phenodeviant surficial shells. This hypothesis is consistent with the observation that extensive shell lags are exposed on the lake floor in areas currently inhabited by the same snails.

A model of biogeographic range fluctuations for particular Paramelania morphs during the Holocene, in concert with lake level fluctuations, can account for the shell assemblages seen on the lake floor today. The complex history of these accumulations suggests that taphonomic admixtures may obscure the interpretation of evolutionary sequences in the lacustrine stratigraphic record.

Survival of North American shelf trilobite families during a mass extinction across the upper boundary of the Upper Cambrian Sunwaptan Stage (=“Ptychaspid Biomere”) cannot be predicted from patterns of turnover among their component species: on the shelf, stratigraphic ranges of species belonging to surviving families do not differ significantly from those of eliminated families. Thus, the sorting of families during the extinctions cannot be explained by simple upward causation from the individual level. However, families that ranged from shelf to slope environments before the extinctions (most of which are pandemic) fared significantly better than those confined to shelf settings (which tend to be endemic to North America), indicating that family survival was influenced by geographic and environmental distribution, a property emergent above the individual level. One factor implicated in clade survival during normal background times, high species richness, did not influence the outcome of mass extinction. The results affirm the importance of a hierarchical approach to the interpretation of macroevolutionary patterns and provide some support to the suggestion that sorting processes operating during mass extinctions differ from those of background times.